Angiosperm Phylogeny

Flowering Plant Systematics

A g
woody, vessels lacking; dioecious; flw T5–8, A∞, G5–8; 1 ovule/carpel; embryo sac 9-nucleate; 1 species (New Caledonia) A mborellales* Amborellaceae
N r
aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A3–∞; embryo sac 4-nucleate
seeds operculate, perisperm; mucilage; alkaloids (no benzylisoquinolines) N ymphaeales * Cabombaceae Hydatellaceae Nymphaeaceae
I a
woody, vessels solitary; flw T>10, A∞, G ca.9; embryo sac 4-nucleate
Austrobaileyales
ethereal oils in spherical idioblasts

T d
e
tiglic acid, aromatic terpenoids Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae
A lvs opposite, interpetiolar stipules; nodes swollen
e flw small T0–3, A1–5, G1, 1 apical ovule/carpel Chloranthales* Chloranthaceae
a woody; pollen uniporate
Canellales
sesquiterpenes

r aromatic terpenoids Canellaceae Winteraceae

l
(pellucid dots)

nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing Aristolochiaceae Piperaceae
y single adaxial prophyll; swollen nodes Piperales Hydnoraceae Saururaceae
woody; lvs opposite; flw with hypanthium, staminodes frequent Calycanthaceae Hernandiaceae Monimiaceae
a often valvate anthers; carpels with 1 ovule; embryo large Laurales Gomortegaceae Lauraceae Siparunaceae
n Magnoliids
benzylisoquinoline alkaloids;

woody; pith septate; lvs two-ranked; ovules with obturator

Annonaceae Eupomatiaceae Magnoliaceae
g features as in
endosperm ruminate Magnoliales Degeneriaceae Himantandraceae Myristicaceae
embryo sac 8-nucleate

“Early Angiosperms”
i infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules

o
ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils
Acorales Acoraceae
s mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules Alismataceae (incl. Limnocharitaceae) Juncaginaceae Ruppiaceae
p flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green Alismatales Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae
Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae
e stem with ring of bundles
r Monocots Ca oxalate
raphides
fr a follicle; East Asia Petrosaviales* Petrosaviaceae
m
Dioscoreales
endosperm often twining vines; lvs often reticulate
atactostele (scattered bundles) nuclear Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae
s no secondary thickening
mostly herbaceous
helobial,
nectaries
ovary often inferior, style short, branched; steroid sapogenins/alkaloids

woody; vessels absent

eustele; nodes
pollen monosulcate
sieve tube plastids with
protein crystals
when present
septal
some woody (with terminally tufted leaves)
flw spadix Pandanales Cyclanthaceae Pandanaceae Velloziaceae
sieve tube plastids often geophytes (bulbs, tubers, rhizomes)
sympodial branching
Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae
with starch grains
lvs simple, persistent, entire
vascular bundles in stem scattered
lvs parallel-veined, entire
flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular
phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins Liliales Colchicaceae Liliaceae Petermanniaceae Smilacaceae
flw parts free, strobilar, no glandular teeth
perfect, P parts varying, flw pentacyclic, P 3-merous, A opp. P often geophytes; anomalous sec. growth Amaryllidaceae ( incl. Agapanthaceae , Alliaceae) Iridaceae
often in threes, filaments narrow
weakly differentiated,
stamen with broad filaments
anthers broadly attached,
septal nectary; single cotyledon
capsule or berry
seed coat obliterated or with phytomelan Asparagales Asparagaceae ( incl. Agavaceae, Hyacinthaceae, Ruscaceae)
Hypoxidaceae Lanariaceae Orchidaceae Tecophilaeaceae
pollen monosulcate adventitious roots
G apocarpous (style short in most) monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate Xanthorrhoeaceae ( incl. Asphodelaceae , Hemerocallidaceae)
nectaries absent
Arecales
intense primary growth, large apical meristem
embryo very small
aromatic terpenoids
infl with spathe; alkaloids Arecaceae
mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae
lvs grassy; flw often anemophilous, minute, chaffy, without nectaries Poales Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae)
flw often irregular; few fertile stamens
Commelinids infl thyrsus of scorpioid cymes
phenylphenalenones Commelinales Commelinaceae Haemodoraceae Pontederiaceae
UV-fluorescing cell walls rhizomatous, large-leafed herbs; pseudostem: central infl
(ferulic/coumaric acids) flw irregular/monosymmetric, septal nectaries
Cannaceae Heliconiaceae Marantaceae Strelitziaceae
silicic acid in leaves
cuticular waxes often in rodlets
aggregated into scallops
G inferior, A often strongly modified/reduced
seeds arillate; silicic acid Zingiberales Costaceae Lowiaceae Musaceae Zingiberaceae
aquatic, herbaceous; lvs whorled, no pellucid dots; vessels lacking; monoecious
flw T9–10, A∞, G1, 1 apical ovule/carpel, pollen inaperturate, pollen tube branched; hydrophilous Ceratophyllales Ceratophyllaceae

Berberidaceae Eupteleaceae Menispermaceae

Ranunculales
lvs often divided; flw whorled, P single or multiple whorls
G apocarpous/paracarpous, superior; berberines Ranunculaceae
Circaeasteraceae Lardizabalaceae Papaveraceae

Sabiales
woody; flw 4–5-merous, nectary: disk, A epipetalous, G syncarpous
pollen colporate; endosperm helobial; triterpenoids Sabiaceae
mostly woody; flw tepals often 4-merous
A epitepalous, connectives sometimes with apical appendage Proteales Nelumbonaceae Platanaceae Proteaceae
E axial
nectary
woody; vessels lacking; idioblasts in cortex; flw tepals missing, A∞
G>5 laterally connate with abaxial nectaries; fr aggregate of follicles Trochodendrales* Trochodendraceae
u common

d mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic

Buxales* Buxaceae Didymelaceae
benzylisoquinolines absent

flw tepals ± uniform or missing; pregnan pseudoalkaloids

i
c unisexual, dioecious; lvs toothed, sec. veins palmate
o
t
flw tepals small to lacking
ellagic acid Gunnerales Gunneraceae Myrothamnaceae
mostly woody; lvs veins proceed to apex & teeth; flw K5–∞, persisting
s
pollen tricolpate
mostly A∞, G mostly slightly connate
seeds often with aril; fr often both ventrally/dorsally dehiscent Dilleniales Dilleniaceae
flw K/C/P opp A lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent
filaments rather narrow Altingiaceae Daphniphyllaceae Hamamelidaceae
Saxifragales
fr mostly dry, dehiscent
anthers basifixed
nodes trilacunar
myricetin, flavonols Cercidiphyllaceae Grossulariaceae Paeoniaceae
stomata anomocytic
often tendrillar vines; lvs often divided and with glandular teeth Crassulaceae Haloragaceae Saxifragaceae
ethereal oils absent

Vitales*
A epipetalous, 2 ovules per carpel; raphides, pearl glands
stipules; berries Vitaceae
nodes
3:3 cork origin deep-seated

Zygophyllales*
endosperm lacking
resinous, lignans/neolignans, harman alkaloids Krameriaceae Zygophyllaceae
ellagic acid tannins

R flw small, G often 3-merous, nectary: intrastaminal disk

Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae)
o
s
seeds often with aril (red-orange)
infl cymose Celastrales Lepidobotryaceae
i lvs often compound, pulvini (sleep movement)
Brunelliaceae Connaraceae Elaeocarpaceae
d Oxalidales
flw A5 or multiple, branched style common
Oxalidaceae
mucilage cells; oxalates Cephalotaceae Cunoniaceae Huaceae
s
F Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae
a lvs margins toothed
flw G often tricarpellate Malpighiales Chrysobalanaceae
Clusiaceae
Hypericaceae
Linaceae
Passifloraceae
Phyllanthaceae
Rhizophoraceae
Salicaceae
b
i flw often “papilionaceous”: wing, standard, keel, mostly G1 Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae
mostly A10; fr a pod; symbiosis with root nodule bacteria
d diverse alkaloids, NP amino acids, lectins (in Fabaceae) Fabales Fabaceae Polygalaceae Quillajaceae Surianaceae
c E s
Barbeyaceae Elaeagnaceae Rosaceae
o u lvs mostly simple with stipules

Rosales
flw K valvate (and hypanthium) persisting
r d N fix
carpels with 1 ovule, stigma dry; dihydroflavonols Cannabaceae Moraceae Ulmaceae
e i embryo large, Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
c endosperm scanty lvs mostly alternate
Anisophyllaceae Coriariaceae Cucurbitaceae
Cucurbitales
flw often unisexual, G mostly inferior
o parietal placentation; cucurbitacins Tetramelaceae
t Begoniaceae Corynocarpaceae Datiscaceae
lvs undivided; flw small, unisexual
s anemophilous, thus T reduced or lacking, G mostly inferior
Betulaceae Fagaceae Myricaceae Rhoipteleaceae
flw pentacyclic
infl spikes or catkins; fr 1-seeded, mostly nuts
ectomycorrhiza; tannins, dihydroflavonols Fagales Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae
parts alternating
lvs with glandular teeth

Geraniales
G connate
flw A obdiplostemonous, nectary on filament
K + C (free)
stems jointed at nodes; ethereal oils, ellagic acid Francoaceae Geraniaceae Ledocarpaceae Melianthaceae
(A polyandrous)
pollen tricolporate
trihydroxyl-flavonoids
lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae)
stipules small (if any); cork deep seated
Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)
flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many
endosperm scanty; scaly bark; flavonols, myricetin Myrtales Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae
woody
Crossosomataceae Stachyuraceae
K + C, stylodia free
hypanthium, nectary disk Crossosomatales Geissolomataceae Staphyleaceae
Strasburgeriaceae
trees; lvs spiral; extrafloral nectaries
M staminate flw: A = and opposite C
bark bitter, anthraquinones Picramniales* Picramniaceae
a mostly woody (silica/silicified)
l Anacardiaceae Meliaceae Rutaceae
v
lvs alternate, odd-pinnately compound
flw often imperfect, intrastaminal disk; ethereal oils Sapindales Burseraceae Nitrariaceae Sapindaceae
Simaroubaceae

i vessel elements: scalariform perforations; mucilage cells
d embryo
lvs margins toothed, stipules cauline
flw small, A = and opposite K, ovules 1-2/carpel Huerteales* Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae
s large
bark fibrous; hairs often stellate Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
endo-
sperm flw K often valvate
scanty petals contorted, A often ∞; mucilage
cyclopropenoid fatty acids, flavones
Malvales Cistaceae
Dipterocarpaceae
Cytinaceae
Neuradaceae
Muntingiaceae
Sphaerosepalaceae
Sarcolaenaceae
Thymelaeaceae
Liverworts
lvs alternate; flw often 4-merous
Mosses often clawed petals, A often many, 2×K Bataceae Caricaceae Limnanthaceae Salvadoraceae
Hornworts
hypogynous (often gynophore); infl racemose
myrosin cells, glucosinolates Brassicales Brassicaceae Cleomaceae Moringaceae Tovariaceae
woody; parasites / semiparasites; without mycorrhiza Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
Lycophytes lvs margin entire; flw A epipetalous
Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae
Ferns
(incl. horsetails)
perianth often simple, valvate, persisting
polyacetylenes, triterpene sapogenins, silicic acid Santalales Loranthaceae Olacaceae Santalaceae (incl. Viscaceae)
Palmferns woody; lvs stomata cyclocytic,

Berberidopsidales*
Ginkgo petiole bundles annular; fr fleshy
calcium oxalate as crystals
Aextoxicaceae Berberidopsidaceae
Ephedra
Welwitschia
mostly herbaceous; without mycorrhiza Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
Seed Gymnosperms Gnetum
Plants Conifers G often unilocular with central placentation, Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
ANITA grade
pollen colpate, surface spiny
betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Caryophyllales [incl. Chenopodiaceae] Droseraceae
Basellaceae Drosophyllaceae
Nyctaginaceae
Phytolaccaceae
Simmondsiaceae
Talinaceae
Magnoliids mostly woody; lvs mostly undivided, hydathode teeth Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
Angiosperms flw often 4-merous, K much smaller than C, persisting,
Monocots

Fabids
intrastaminal disk, G inferior; fr drupaceous
diverse iridoids Cornales Cornaceae
Curtisiaceae
Grubbiaceae
Hydrangeaceae
Loasaceae
Nyssaceae
Rosids Malvids lvs teeth, theoid; nodes unilacunar Actinidiaceae Ericaceae Polemoniaceae Sarraceniaceae
flw 5-merous, pentacyclic
Balsaminaceae Fouquieriaceae Primulaceae Styracaceae
Lamiids nonhydrolyzable tannins, ellagic acid, hydroquinones
Ericales Clethraceae Lecythidaceae Roridulaceae Theaceae
Asterids Campanulids
woody; dioecious Ebenaceae Myrsinaceae§ Sapotaceae Theophrastaceae
flw small, C valvate, G unilocular; fruit indehiscent

A
iridoids (aucubin), gutta
Garryales Eucommiaceae Garryaceae (incl. Aucubaceae)
lvs opposite, colleters
Theodor C. H. Cole, Dipl. Biol. s Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Institute of Pharmacy and Molecular Biotechnology t late
sympetaly
flw corolla convolute in bud
indole alkaloids; iridoids Gentianales Gentianaceae Gelsemiaceae Rubiaceae
Heidelberg University e
Im Neuenheimer Feld 364 lvs spiral, simple; nodes unilacunar
r Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)
D-69120 Heidelberg, Germany
i L flw petals plicate; K persisting
diverse alkaloids, no iridoids Solanales Hydroleaceae Montiniaceae Sphenocleaceae
d a
m lvs opposite; nodes 1:1; flw mostly monosymmetric Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
s A=C,
ii
A often 2(+2); ethereal oils in gland-headed hairs
Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
Lamiales
epipetalous 6-oxygenated flavones, rosmarinic acid,
mostly sympetalous polyandry d oligosaccharides: cornoside, verbascoside (acetoside) Byblidaceae Martyniaceae Pedaliaceae Stilbaceae
s
rare,
nectary gynoecial
ovules unitegmic G(2)
lvs roughly hairy; nodes unilacunar
Gesneriaceae Hydrostachyaceae Oleaceae Phrymaceae Verbenaceae
endosperm cellular
Boraginales
infl scorpioid; mostly 4 ovules
iridoids common
infl
cymose isokestose, higher inulins, pyrrolizidine alkaloids
Boraginaceae Codonaceae Wellstediaceae Hydrophyllaceae
ellagic acid Ehretiaceae (+ Lennoaceae) Cordiaceae Heliotropiaceae (Namaceae)
lacking woody; lvs serrate;
Prof. Dr. Hartmut H. Hilger
Dahlem Centre of Plant Sciences (DCPS) early
flw 1-2 ovules/carpel, C± free, K slightly connate
fr drupe with broad stigma Aquifoliales Aquifoliaceae Cardiopteridaceae Stemonuraceae
sympetaly
Institute of Biology – Plant Morphology and Systematics flw small flw often monosymmetric
Asteraceae Goodeniaceae Pentaphragmataceae
Freie Universität Berlin plunger pollination (long style)
Asterales
embryo
Altensteinstr. 6, D-14195 Berlin, Germany short nodes trilacunar Calyceraceae Menyanthaceae Rousseaceae
inulin, sesquiterpenes, secoiridoids
Campanulaceae (incl. Lobeliaceae) Stylidiaceae
________________________________________________________________________________________
- hypothetical tree based on molecular phylogenetic data (Feb. 2014); branch lengths deliberate, not expressing actual time scale
C mostly woody; infl racemose, C free
anthers basifixed, nectary disc Escalloniales* Escalloniaceae
- position of many characters on tree unclear; if a character is marked as being a potential synapomorphy at a node/for a clade, a
this does not mean that all members of that clade possess that character; * orders added as of APG III (2009)
- this poster depicts only the largest and most important of the currently accepted approx. 450 families (according to APweb 2014);
m woody, evergreen; nodes 1:1
flw polysymmetric, anthers basifixed Bruniales* Bruniaceae Columelliaceae (incl. Desfontainia)
p
G inferior

for family characteristics see: Kubitzki K, ed. (1990 ff).

- References: APG III (2009); Judd W et al. (2007); Simpson M (2010); Soltis DE et al. (2005/2011/2014); a woody (except Apiaceae)
lvs often divided; nodes usu. multilacunar Apiaceae Griseliniaceae Pennantiaceae
Stevens PF (2014) APweb – www.mobot.org/MOBOT/research/APweb; Watson/Dallwitz (2009) delta-intkey.com/angio/
§ Myrsinaceae and Theophrastaceae again included in Primulaceae by APG III
n
u
infl mostly umbel; drupe or schizocarp
(Apiaceae: mericarp/carpophore)
A piales Araliaceae Myodocarpaceae Pittosporaceae
1–few-seeded

l
fr usually

woody; infl racemose, flw 4-merous,

Thanks to Christoph Dobeš, Ray F. Evert, Marc Gottschling, Richard G. Olmstead, Peter H. Raven, Douglas E. Soltis, Peter F. Stevens, Maximilian Weigend, Michael Wink
i filaments stout, capsule septicidal Paracryphiales* Paracryphiaceae
Angiosperm Tracheophte Bryophte d lvs opposite, often basally connate
Phylogeny Phylogeny Phylogeny s nodes 3:3; buds with scales
Adoxaceae Diervillaceae Linnaeaceae
Poster Poster Poster
flw often monosymmetric
K persistent in fruit; secoiridoids Dipsacales Caprifoliaceae Dipsacaceae Morinaceae
Valerianaceae